Sodium (Na+) homeostasis and salt tolerance of plants

• Salinity adversely affects crop yield and agricultural expansion.
• Ion transport systems facilitate Na+ homeostasis and salt tolerance.
• HKT1 is a model for translation of basic knowledge to crop salt tolerance.
• Halophytes are sources for unique alleles and loci of ion homeostasis determinants.

Soil and water salinity substantially constrain crop and biomass production. Research over the last two plus decades, facilitated by advances in molecular genetics and biotechnology, and with genetic model systems, has identified genes involved in salt acclimation or adaptation and linked these to critical mechanisms and processes. A case in point is present understanding of critical transport determinants that facilitate intra- and intercellular Na+ homeostasis of plants in saline environments predominated by NaCl. Pumps in the plasma membrane (H+-ATPase), and the tonoplast (H+-ATPase) and H+ pyrophosphatases (AVP1) generate proton electrochemical gradients necessary to energize Na+ efflux to the apoplast and influx into vacuoles, respectively. The plasma membrane Na+/H+ antiporter SOS1 is responsible for apoplastic efflux, and NHX type Na+/H+ antiporters for vacuolar and endosomal compartmentalization. Ca2+ext reduces passive intracellular Na+ influx cells by decreasing Na+ transport through high affinity K+ uptake systems and what are presumed to be nonselective cation channels, and activating, through the SOS signal pathway, the SOS1 plasma membrane Na+/H+ antiporter. Moreover, there is greater understanding about how cellular transport systems functionally integrate to facilitate tissue and organismal Na+ homeostasis. Notable in this process are HKT1 Na+ transporters, which regulate Na+ loading into the root xylem, limiting flux to and accumulation in the shoot. This review will summarize ion transport systems that facilitate plant Na+ homeostasis. Halophyte and glycophyte salinity responses and transport determinant function are compared and contrasted. The potential of halophytes as genetic resources for unique alleles or loci of transport protein genes and transcriptional and post-transcriptional regulation of transport protein function are discussed in the context of crop salt tolerance.